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Zoology. Proceedings of the Fiftieth Anniversary Meeting of the Society for Experimental Biology PDF

539 Pages·1976·13.26 MB·English
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PERSPECTIVES IN EXPERIMENTAL BIOLOGY Volume 1 Zoology Proceedings of the Fiftieth Anniversary Meeting of the Society for Experimental Biology Edited by P. SPENCER DAVIES With the editorial assistance of A. PUNT, G. M. HUGHES, S. H. P. MADDRELL, J. E. TREHERNE, D. B. SATTELLE, A. P. M. LOCKWOOD, J. D. ROBERTSON and E. R. TRUEMAN P E R G A M ON PRESS OXFORD · NEW YORK · TORONTO · SYDNEY PARIS · BRAUNSCHWEIG U. Κ. Pergamon Press Ltd., Headington Hill Hall, Oxford 0X3 OBW, England U. S. A. Pergamon Press Inc., Maxwell House, Fairview Park, Elmsford, New York 10523, U.S.A. CANADA Pergamon of Canada, Ltd., 207 Queen's Quay West, Toronto 1, Canada AUSTRALIA Pergamon Press (Aust.) Pty. Ltd., 19a Boundary Street, Rushcutters Bay, N.S. W. 2011, Australia FRANCE Pergamon Press SARL, 24 rue des Ecoles, 75240 Paris, Cedex 05, France WESTGERMANY Pergamon Press GMbH, 3300 Braunschweig, Postfach 2923, Burgplatz 1, West Germany Copyright © Pergamon Press 1976 All Rights Reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means: electronic, electrostatic, magnetic tape, mechanical, photocopying, recording or otherwise, without permission in writing from the publishers First edition 1976 Library of Congress Catalog Card No. 75-34712 Printed in Great Britain by A. Wheaton & Co., Exeter ISBN 0 08 018767 6 (Volume 1) 0 08 019868 6 (Volume 2) 0 08 019939 9 (2 Volume set) LIST OF CONTRIBUTORS ALEXANDER, R. MCN., Department of Pure and Applied Zoology, University of Leeds. ANDERSEN, SVEND OLAV, August Krogh Institute, University of Copenhagen, Denmark. ANWYL, R., Department of Zoology, The University, Glasgow, U.K. ATKINSON, R. J. Α., Department of Marine Biology, University of Liverpool, Port Erin, Isle of Man, U.K. BATESON, P. P. G.,' University of Cambridge, Sub-Department of Animal Behaviour, Madingley, Cambridge, U.K. ΒΕΝΝΕΤ-CLARK, H. C, Department of Zoology, University of Edinburgh, Edinburgh, U.K. BITENSKY, LUCILLE, Division of Cellular Biology, The Mathilda and Terence Kennedy Institute of Rheumatology, Bute Gardens, London, W6 7DW, U.K. BOWLER, K., Department of Zoology, University of Durham, Durham, U.K. BROWN, H. MACK, Department of Physiology, University of Utah Medical Centre, Salt Lake City, Utah, U.S.A. BURROWS, MALCOLM, Department of Zoology, University of Oxford, U.K. BURTON, R. F., Institute of Physiology, University of Glasgow, Glasgow, U.K. CHAYEN, J., Division of Cellular Biology, The Mathilda and Terence Kennedy Institute of Rheumatology, Bute Gardens, London, W6 7DW, U.K. CLARK, R. B., Department of Zoology, University of Newcastle upon Tyne, U.K. CRISP, D. J., N.E.R.C. Unit of Marine Invertebrate Biology, Marine Science Laboratories, Menai Bridge, U.K. CROGHAN, P. C, School of Biological Sciences, University of East Anglia, U.K. DAVIES, P., Division of Cell Pathology, M.R.C. Clinical Research Centre, Watford Road, Harrow, Middx., U.K. DORSETT, D. Α., Marine Science Laboratories, University College of North Wales, Bangor, U.K. DUNCAN, C. J., Department of Zoology, University of Liverpool, U.K. FLETCHER, C. R., Department of Pure and Applied Zoology, University of Leeds, U.K. GOLDSWORTHY, G. J., Department of Zoology, University of Hull, U.K. ix χ LIST OF CONTRIBUTORS GUPTA, BRU L., Department of Zoology, Downing Street, Cambridge, U.K. HUDDART, HENRY, Department of Biological Sciences, Lancaster University, Lancaster, U.K. HUGHES, G. M., Research Unit for Comparative Animal Respiration, Bristol University, Bristol, U.K. HUTCHISON, J. B., MRC Unit on the Development and Integration of Behaviour, University Sub-Department, Madingley, Cambridge, U.K. JOHANSEN, KJELL, Department of Zoophysiology, University of Aarhus, DK-8000 Aarhus C, Denmark. LOCKWOOD, A. P. M., Department of Oceanography, University of Southampton, U.K. MADDRELL, S. H. P., A.R.C. Unit of Invertebrate Chemistry and Physiology, Cambridge, U.K. MAETZ, J., Departement de biologie du C.E.A., Station Zoologique, 06230 Villefranche sur Mer, France MEECH, ROBERT W., A.R.C. Unit of Invertebrate Chemistry and Physiology, Depart- ment of Zoology, Cambridge, U.K. MILLER, PETER L., Department of Zoology, Oxford, U.K. MILLS, PAULINE S., Department of Zoology, Oxford, U.K. NAYLOR, E., Department of Marine Biology, University of Liverpool, Port Erin, Isle of Man, U.K. PAYAN, P., Departement de biologie du C.E.A., Station Zoologique, 06230 Villefranche sur Mer, France. PHILLIPS, J. E., Department of Zoology, University of British Columbia, Vancouver 8, B.C., Canada. PICHON, YVES, Unite de physiologie de l'Insecte, Laboratoire de Neurobiologie Cellulaire, Centre National de la Recherche Scientifique, 91 190 Gif sur Yvette, France. PIDDINGTON, R. W., A.R.C. Unit of Invertebrate Chemistry and Physiology, Department of Zoology, University of Cambridge, Downing Street, Cambridge, U. K. POTTS, W. T. W., Department of Biological Sciences, University of Lancaster, U.K. PRINCE, W. T., A.R.C. Unit of Invertebrate Chemistry & Physiology, Department of Zoology, University of Cambridge, U.K. RENZIS, G. DE, Departement de biologie du C.E.A., Station Zoologique, 06230 Ville- franche sur Mer, France. SCHOFFENIELS, E., Laboratory of General and Comparative Biochemistry, University of Liege, 17 Place Delcour, B-4000 Liege, Belgium. LIST OF CONTRIBUTORS xi SHELTON, G., School of Biological Sciences, University of East Anglia, Norwich, U.K. SLATER, T. F., Biochemistry Department, Brunei University, Uxbridge, Middlesex, U.K. SLEIGH, MICHAEL Α., Department of Biology, University of Southampton, U.K. SMITH, C. L. Department of Zoology, University of Liverpool, U.K. SMITH, R. I., Department of Zoology, University of California, Berkeley, California, 94720, U.S.A. SUTCLIFFE, D. W., F.B.A., Windermere, U.K. TREHERNE, J. E., A.R.C. Unit of Invertebrate Chemistry and Physiology, Department of Zoology, Cambridge, U.K. TRUEMAN, E. R., Zoology Department, University of Manchester, U.K. USHERWOOD, P. N. R., Department of Zoology, University of Nottingham, Nottingham, U.K. WAREHAM, A. C, Department of Physiology, Medical School, University of Manchester, Manchester, U.K. WEBB, J. E., Department of Zoology, Westfield College, London, U.K. WEBER, ROY E., Department of Zoophysiology, University of Aarhus, DK-8000 Aarhus C, Denmark. WEIS-FOGH, TORKEL, Department of Zoology, Downing Street, Cambridge, U.K. WELLS, G. P., Department of Zoology, University College, Go wer Street, London, U.K. WELLS, M. J., Zoology Department, University of Cambridge, U.K. PREFACE The Fiftieth Anniversary of the inauguration of the Society for Experimental Biology was celebrated by a special meeting held at the University of Cambridge from July 16th to 19th, 1974. Prominent members of the Society were each invited to present a paper on their own specialized field of research, providing a historical review, a statement of current thought in that area and, if possible, a predictive element. In the belief that the material thus presented would be of considerable use to other research workers, the speakers were invited to contribute their papers to these two volumes. The chapters therefore are each a personal affirmation, and reflect the approaches, opinions and styles of their individual authors. For convenience, the papers have been collected into one zoological and one botanical volume. Within each volume papers on a particular discipline have been grouped together, although these are not necessarily the groupings that were used when the papers were read at the meeting. This perhaps underlines the difficulty of classification in such a diversified field as is here presented. However, this diversity gives an indication of the breadth of the Society's interests and of the topics which tend to appear most frequently in its programmes. The diversity too, emphasizes the strength and aims of the Society; to provide a forum for research workers in both animal and plant sciences to discuss current research, to present work which is often still in progress and incomplete, and above all, to encourage the cross-fertilization of ideas between specialized disciplines. It is hoped that the material assembled in these two volumes will provide for young research workers a perspective of the field of experimental biology and for under- graduates a source of reference to a wide range of biological topics. Several invited speakers were unable to provide a manuscript and we wish to record our thanks to them for their contribution to the Anniversary meeting. We are also indebted to those distinguished members of the Society who acted as chairmen to the sessions of the meeting. They were T. Weis-Fogh, R. Brown, G. M. Hughes, A. Punt, P. W. Brian, Sir Vincent Wigglesworth, J. Chayen, R. D. Keynes, F. R. Whatley, E. W. Simon, G. E. Fogg, J. Heslop-Harrison, J. W. S. Pringle, Sir Rutherford Robertson, J. S. Kennedy, L. C. Beadle, D. A. Boulter, G. P. Wells, J. D. Robertson and Helen K. Porter. Finally, particular thanks are due to the local organizing team: Helen Skaer, D. A. Hanke, C. G. Gill and the able Local Secretary, D. B. Sattelle. Ν. SUNDERLAND, P. SPENCER DAVIES, John Innes Institute, Department of Zoology, Colney Lane, The University, Norwich, U.K. Glasgow, U.K. xiii T HE EARLY DAYS OF T HE S.E.B. G. P. WELLS Department of Zoology, University College* Gower Street, London, U.K. THE Society for Experimental Biology was called into being in December 1923, to support a Journal, then known as the British Journal of Experimental Biology. The Society has always had botanists among its members, but in its early years it was dominated by zoologists. The first published Part of the Journal contained six papers, all zoological. The programme of the Conference at which the Society was set up, included twenty papers, only three of which were botanical. Later, the botanical side grew very rapidly until it came to match the zoological, but in the beginning it was the younger zoologists who most urgently needed a Society, and a Journal, to cater for the experimental aspects of their subject. The reason is that a great darkness had settled on the majority of British zoologists in the early years of this century. They became obsessed by comparative anatomy and descriptive embryology, and by the possible evolutionary relationships of the animals whose corpses they studied. Lancelot Hogben (1970) has traced for us the slow reawakening of experimental zoology in Britain in the years before World War I, and its more vigorous expansion in the early twenties. But even in the mid-twenties, there were still many centres of learning in which the idea that zoology could be an experimental science had hardly dawned. A fair way of showing what the word 'Zoology' then meant, to most of the people who used it, is to quote the syllabus of the B.Sc. courses in Zoology in London in 1924. London, besides being a great university, was, then more than now, a great examining body, controlling, through its External Degree examinations, a number of important teaching centres that have since become Universities in their own right. The syllabus that I am about to sketch out determined the form and content of the subject for hundreds of students, many of whom were destined to become teachers in their turn, so its influence extended far and wide. Suppose then that, fifty years ago, you were interested in zoology, and set out to get a London B.Sc. degree. You could take a General or a Special B.Sc. Either would keep you busy for at least 2 years after you had passed the Intermediate Examination. The General Degree Course consisted of three science subjects of equal weight, one of which, in your case, would be zoology. The Special Degree Course would consist mainly of intensive teaching in zoology, with a lesser amount of one other science subject : its syllabus was an elaboration of that for the General Course. Here is what you must master, if you wanted to pass in zoology at the General level in 1924: The main facts and principles of Zoology as exemplified by a study of the characteristic structure and development of typical members of all the larger groups of the animal kingdom, with specia 1 2 G. P. WELLS reference to those contained in the syllabus of practical work which follows; the chief facts and the theories of heredity and evolution; the classification of the animal kingdom and the distinguishing characters of the larger groups. Then follows the list of types which is to form the basis of the practical course. It consists of 258 named animals. Slightly more than half of these are distinguished by asterisks; this 'signifies external form or demonstration specimen only'. The remaining 121 you must know pretty well. The types range from Amoeba to Man, but Man has an asterisk. In addition to this, you must know twenty-three named larval forms, and the early stages in the development of Amphioxus, frog, chick and rabbit. This ends the syllabus. If you aspired to Special Honours in zoology you had to know all this, together with a more extended study of the more modern aspects of zoological work with special reference to (1) Evolution and its evidences, (2) Bionomics, (3) Heredity (Genetics), (4) Experimental Morphology and Embryology, (5) the meaning of sex and the mechanism of its determination. You also studied a limited topic, known as a Special Subject, 'as far as possible from original publications and actual material'. This topic you chose in consultation with your teachers. Notice that these courses contained no physiology, no behaviour, very little ecology unless there was some in your Special Subject. Even those parts of the prescribed course which are essentially experimental—genetics and experimental embryology—were usually taught by lectures alone. In his practical examination, the candidate was required to do three things : to dissect animals, to make sections and other permanent preparations, and, confronted by rows of specimens, to indentify, draw and classify them, 'giving your reasons'. Meanwhile, the Botany students had a much better time. Their syllabus included substantial amounts of physiology and ecology; their material lived; they made experi- ments in the teaching laboratory and in the examination room. But animal physiology stood apart from zoology and was taught as a separate subject with a strong medical bias. In those days it was generally restricted to the mammal, coupled with the central nervous system, heart and hind legs of the frog. This was the situation from which the experimentally-minded zoologists were breaking away. One of the most active centres of the new movement was in Cambridge, and its leader was James Gray. Though not yet Professor, he became, in Hogben's words, 'the mentor of a new generation of Cambridge zoologists'. Others, too, were experimenting in the zoology department—notably Munro Fox and J. T. Saunders—and experimental questions began to appear in both written and practical examination papers. Comparative physiology was also attracting attention elsewhere—in the Department of Physiology for example, where Joseph Barcroft was extending his studies on haemoglobin to invertebrates. One can learn something of the extent and variety of the movement by turning over the pages of the first volume of the Biological Proceedings of the Cambridge Philosophical Society, a volume which covered the 2 years from 1923 to 1925. It contains contributions in other fields—in palaentology and botany, for example—but nearly two thirds of its twenty-nine papers are about comparative physiology and experimental THE EARLY DAYS OF THE S.E.B. 3 zoology. This, of course, is not because the followers of these lines of enquiry were more numerous, or more productive, than their colleagues. It is because the workers in other fields had other journals in which they could readily publish. It was one of the chief reasons why the Biological Proceedings was started, to provide an outlet for the experi- mentally-minded zoologists. However, it was not in Cambridge that our Society began. We were conceived in Edinburgh and born in London, at Birkbeck College. On the other hand, the British Journal of Experimental Biology, for whose support we were created, was born in Edin- burgh but conceived in London—more precisely, in the head of Hogben, on whose first-hand account (1970) I have based my narrative of these events. Lancelot Hogben, a man of tremendous energy and breadth of vision, was a Cam- bridge graduate who became, for a few years, a lecturer at the Royal College of Science in South Kensington. Here he had what he has modestly described as 'a by no means epoch-making flirtation with genetical cytology'—a flirtation that resulted in six papers and an invitation to move to Edinburgh. The invitation came from Frank Crew, an Edinburgh graduate of medicine with a strong interest in genetics, who had abandoned medical practice after demobilization from World War I to teach zoology in his old University. Presently there was set up in Edinburgh a new institute—the Animal Breeding Research Department—and Crew became its first director in 1921. He was anxious to have a cytologist around. So Hogben was invited to go north, and joined Crew as his second-in-command. By that time he was turning from cytology to endocrinology and the physiology of colour change, and he combined his position at the Animal Breeding Research Station with a part-time lectureship in Comparative Physiology in the Department of Zoology. While still at the Royal College of Science, Hogben had begun to think of a biological journal for the publication of experimental research. When he reached Edinburgh, he found Crew's Department, in his own words 'temporarily housed by Crew's inspired opportunism in a dilapidated and abandoned Infirmary in a back street of Old Edin- burgh'. But, in spite of its shabbiness, many distinguished biologists came for working visits. Among them were Julian Huxley and J. B. S. Haldane, and these two completed the quartet of, in Hogben's phrase, 'the Founding Fathers of the S.E.B.'. With the other three, Hogben discussed his idea for a new Journal. Crew at once declared that he had in hand enough cash from compensation for war wounds to finance the initial project, and that he would be delighted to use it in this way. For other reasons, and not because of his generous offer, the other three prevailed on Crew to become the first Managing Editor and a supporting Editorial Board of eleven names was organized, including a Professor of Botany, Ruggles Gates. Crew persuaded Oliver & Boyd to print and publish the new journal in Edinburgh. Hogben undertook the circularization of libraries, University departments, and Institutes. Finally, in October 1923, the first Part of the British Journal of Experimental Biology saw the light. Crew was then 35 years old, Huxley 36, Haldane 31 and Hogben 28. It seemed to the Founding Fathers that a Society was needed, as a secure base for the B.J.E.B. If one belonged to such Societies as the Physiological, the Biochemical or the Linnaean in those days, one paid a handsome subscription and automatically received the Society's journal, which could thus rely on financial support. Moreover, one generally 4 G. P. WELLS sent one's manuscripts to one's journal. To provide the Β J.E.B, with these advantages, the Founding Fathers (overcoming some resistance from other members of the Editorial Board) announced a Conference to inaugurate the new Society, to take place at Birkbeck College in the Christmas vacation, 1923. The invitations went to University departments of botany, zoology and physiology and to research institutes, and in the event the attendance exceeded the most extravagant hopes of the promoters. That inaugural conference, as anybody who was there will remember, was a tremend- ously exciting occasion. From then on, the success of the S.E.B, was assured—except in one respect. It had been invented to give support to the B.J.E.B., and this it failed to do. Paradoxically perhaps, its failure was due to the peculiar virtues which ensured its success as a Society. These virtues were two. In the first place, the great majority of its members were young. In the second place, its interests were not restricted to a single specialized field. It brought together experimental biologists from a range of different specializations, to bring about, in Hogben's phrase, 'the cross-fertilization between widely separated compartments of biological enquiry'. But the young were not opulent; the members of specialist societies who joined us were already paying subscriptions elsewhere; and so it was decided that one could belong to the S.E.B, for 12s. 6d. per annum. If, in addition, one took the journal, the subscription was £2. But it turned out that only a minority of members chose the latter option. So what was to be done, to ensure the financial stability of the B.J.E.B. ? Its difficulties were increased by the appearance, 2 months before its first issue came out, of another journal about which I have already spoken—the Biological Proceedings of the Cambridge Philosophical Society. Experimental zoology certainly needed an outlet, but not, at that stage, two. So there were conversations, and meetings, and a solution was found. The B.J.E.B. had included from the first, besides papers reporting on individual researches, a certain proportion in which the writers reviewed recent progress in their fields. This function was taken over by the Cambridge journal, which became Biological Reviews, and under the editorship of Munro Fox, at once achieved international importance. Papers on particular experimental researches became the business of the B.J.E.B. But still the financial position of the B.J.E.B. was insecure, until the problem was solved by the intervention of G. P. Bidder. He was a Cambridge zoologist with sub- stantial private means and business interests, an expert on sponges, a familiar figure at Naples and at Plymouth, who had assisted progressive developments in the biological sciences more often, I suspect, than is generally known. His contribution to our story was the invention of the Company of Biologists Limited, as a stable financial foundation for the journal. He induced forty or fifty biologists to take shares in the Company, and himself guaranteed any overdraft up to a sufficient limit. With the funds so raised, the B.J.E.B. was bought from Oliver & Boyd, and thereafter was published for the Company by the Cambridge University Press. The Company appoints the Editor of the journal, but does not meddle with what he does. Wisely, they persuaded James Gray to undertake the Editorship. Their business is managed by a Board of Directors, consisting at first of Crew, Huxley, Hogben, Saunders and Bidder, who was also the Secretary. Bidder resigned from the Secretaryship after 3 years, to be replaced by Saunders. The word British' was later dropped from the title of the Journal, because of a short-lived plan to

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